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Stewart, K.W., B.P. Stark, & L.L. Serpa. 2013. Larvae of the two North American species of Calileuctra (Plecoptera:

Leuctridae). Illiesia, 9(01):1-13. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia09-01.pdf

urn:lsid:zoobank.org:pub:41507049-00D1-4738-8EF1-28EC83CDA1D2

Illiesia – http://www2.pms-lj.si/illiesia/ Volume 9 – Number 1 –

LARVAE OF THE TWO NORTH AMERICAN SPECIES OF CALILEUCTRA

(PLECOPTERA: LEUCTRIDAE)

†Kenneth W. Stewart1, Bill P. Stark2 and Larry L. Serpa3

1 Department of Biological Sciences, P.O. Box 305220, University of North Texas, Denton, Texas 76203, U.S.A.

E-mail: [email protected]

2 Department of Biology, Box 4045, Mississippi College, Clinton, Mississippi 39058, U.S.A.


3 The Nature Conservancy, 201 Mission Street, San Francisco, California 94105 U.S.A.


ABSTRACT

Comparative larval descriptions and an updated generic diagnosis are provided for the two rare Nearctic

species of the leuctrid genus Calileuctra Shepard & Baumann. The larva of the generotype C. ephemera Shepard

& Baumann,was previously described, in part, from a single larva that was illustrated and subsequently lost,

and the larval exuvium of a single reared male paratype. Larvae of C. dobryi Shepard & Baumann,were

previously described in part from three individuals. Comparative descriptions utilize additional associated

larvae, and SEM and light microscope study, providing additional characters. A key to the larval species is

provided and differences in the molar structure of the right mandible of Leuctridae and Capniidae are

discussed.

Keywords: Calileuctra, Plecoptera, Leuctridae, Larval descriptions

INTRODUCTION

The leuctrid genus Calileuctra Shepard &

Baumann 1995, currently includes the two rare

species C. ephemera Shepard & Baumann 1995, and C.

dobryi Shepard & Baumann 1995, both known only

from California. These species inhabit mostly

intermittent headwater stream tributaries, including

their type localities and additional sites. Adults are

present from December to March, depending on

seasonal precipitation; largest numbers have been

collected in February. Based on the intermittent

nature of the type localities and other known

collection sites, and our unsuccessful collection

attempts in drought years, the life cycle is probably a

fast univoltine type, or fast semivoltine in successive

drought years, similar to species of Zealeuctra

(Snellen & Stewart 1979); this, and possible

hyporheal occurrence of larvae until just before

emergence, have made it difficult to get the timing

right for collecting good numbers of mature larvae

for study.

Shepard & Baumann (1995) gave a written

Stewart, K.W., B.P. Stark, & L.L. Serpa. 2013. Larvae of the two North American species of Calileuctra (Plecoptera: Leuctridae).

Illiesia, 9(01):1-13. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia09-01.pdf


description of the single known larva of C. ephemera

that they collected with the types. That larva,

indicated as deposited in the California Academy of

Sciences Museum, was subsequently lost before 2000.

A habitus of it, drawn by Jean Stanger-Leavitt prior

to its loss, was used with permission of R.W.

Baumann for the partial description and illustration

of the larva by Stewart & Stark (2002). Attempts by

us to collect more larvae at the Capell Creek type

locality in February, 2000, and early March, 2012,

when it had only mud substrate or recent rainwater

pools, were unsuccessful. Author (L. Serpa) has

subsequently collected numerous adults and 15

larvae in sand-cobble substrates of pools from

intermittent headwater tributaries, that flow

variously, for up to nine months, located in oak-

dominated areas of Douglas fir (Pseudotsuga menziesii

(Mirb.) Franco) and Redwood (Sequoia sempervirens

(D.Don) Endl. forests in Mendocino County.

The four larvae of C. dobryi, studied and partially

described by Stewart & Drake (2007), and an

additional exuvium were available for this

comparative study. Previous SEM’s of mouthparts

they described, of one larva, were reexamined for

different views, along with further SEM’s of one

other larva, leaving two remaining as vouchers for

possible future morphological study. Several

attempts to collect more larvae of C. dobryi have been

unsuccessful. In mid-February, 2000, author (K.

Stewart), accompanied by Keith Dobry, for whom the

species is named, attempted for two days to collect

larvae and adults at the Ellsmere Canyon type

locality, and other known intermittent area streams

of sagebrush-oak woodlands in Los Angeles County,

to no avail. A large population of adults, the 4 larvae

and the exuvium were subsequently found in

Silverado Canyon Creek in Orange County by K.W.

Stewart and E.F. Drake from 2004-2006. The

substrates of slower-flow habitats, where C. ephemera

larvae have recently been found, may not have been

adequately sampled for larvae at this site.

MATERIALS AND METHODS

Larvae were examined with light and scanning

microscopy. Drawings were prepared using an

Olympus SZH 10 or Wild M5 dissecting microscope

with drawing attachment. Comparative SEM’s were

taken of specific body structures using an

Amray1810D scanning electron microscope. The

specimens studied are deposited in the B.P. Stark

Collection, Mississippi College, Clinton, Mississippi,

and L. Serpa personal collection. Because of the small

sample size of larvae of these two rare species

available for study, descriptions and the key do not

address potential individual or geographic variation

of characters. We have confidence in the field

associations of larvae because of rarity of the species

and that no congeners were found in any of the

collection sites.

RESULTS AND DISCUSSION

Generic Diagnosis of Calileuctra

Following is an expanded and updated generic

description of Calileuctra larvae from that of Stewart

& Stark (2002) that was based on the partial

description of C. ephemera larvae. The originally

proposed combination of diagnostic Calileuctra

characters included: 1) Ab1-7 divided by a pleural

fold, (this character is shared only with the leuctrid

genus Megaleuctra Neave, which is much larger and

more setose, and with Perlomyia Banks, which has

fused, terminally hairless paraprocts and a single

stem of the mesosternal Y-ridge), 2) right mandible

molar area with 8-10 stalked teeth in side view,

(originally described by Stark & Stewart (2002) as a

“scraping ridge”), and 3) a double stem of the

mesosternal Y-ridge, (this character shared only with

Paraleuctra Hanson and Pomoleuctra Stark & Kyzar,

but both have much longer terminal cercomere

hairs). These original characters are all supported by

this study and the generic description, given below,

is updated with revised and newly discovered

characters, measurements, and illustrations. The

further SEM study, with views into the right

mandibular cup of C. dobryi, show stalked-tooth

ridges similar to those described for C. ephemera by

Shepard & Baumann (1995), and comparative side

views of the cup of other leuctrid and capniid genera

(Stewart & Stark 2002), suggest that the mandibular

ridges found in leuctrid genera may be an additional

character used in the difficult separation of larvae

from these families.

Larval morphology. Body length ♂ 5-7mm, ♀ 6-

8mm, light colored body with little pigmentation and

generally few hairs (Figs. 1, 21). Antennae with 60-64




Fig. 1. Calileuctra ephemera pharate larval male habitus. Scale line = 2 mm.

segments, each with apical circlet of very short hairs

or sensillae. Mouthparts Type 1 (herbivorous-

detritivorous; Stewart & Stark 2002). Labrum and

clypeus with numerous setae. Lacinia triangular and

palmate, with 2 large, round-tipped apical teeth,

dorsal and ventral rows of sharp spines, and a

scalloped palm surface (Figs. 7, 8). Right mandible

with short, rounded-tipped apical cusps and molar

cup in side view with 8-10 short, stalked teeth (Figs.

11, 26); these teeth are manifested as curved ridges

extending well onto the cup surface (Fig. 27). Left

mandible with similar apical cusps; its molar cup

with about 12 long, outer, finger-like teeth (Fig. 12)

and inner surface with raised, transverse ridges (Fig.

13). Pronotum quadrangular, with no marginal setae

and no distinct pattern (Figs. 1, 21). Mesosternal Y-

ridge with a double stem, arms reaching to posterior

corners of furcal pits, and a transverse ridge

connecting anterior corners of furcal pits (Fig. 2).

Wingpads with few, or no, hairs, 3 or more times

longer than wide and with their longitudinal axes

nearly parallel to body axis; hind wings slightly

shorter than fore wings (Figs. 1, 21). Legs slightly

increasing in size posteriorly; femora and tibiae with

appressed, fine clothing hairs, scattered, short thick

hairs (Fig. 14), and tarsal segments 1, 2 with a double

row of short thick setae (Fig. 15). Abdominal terga

without a posterior fringe of setae and variable

1




Figs. 2-6. Calileuctra ephemera larval characters. 2. Male mesosternum. 3. Pharate male terminalia, dorsal.

4. Pharate male terminalia, ventral. 5. Female terminalia, ventral. 6. Cercus, dorsal. Scale line = 1mm,

arrows = developing male genitalic features.

surface setation between species; pleural fold on

segments 1-7 (Figs. 5, 23). Developing male genitalia

evident dorsally and ventrally (Figs. 1, 3, 4) just prior

to emergence. Paraprocts only incompletely fused at

base (Fig. 5), each with a pair of terminal hairs (Figs.

1, 3, 4, 5). Cercal segments 20-22, each with apical

circlet of more than 12 stiff setae, half as long as, or

longer than, following segment (Figs. 6, 20, 24).

Species Accounts of Larvae

The following accounts include: 1) known

distribution, 2) larvae, exuviae and adult material

examined, and 3) description of characters not

addressed in the generic diagnosis above, or that

offer specific diagnosis, reinforcement or correction

of original character descriptions of C. ephemera by

Shepard & Baumann (1995) and Stewart & Stark

(2002), and for C. dobryi by Stewart & Drake (2007).

Descriptions and illustrations are based on typical,

late instar individuals or exuviae from the few

populations and individuals that were successfully

field associated; therefore, they do not address

possible variation of characters.

2 3

4 5

6




Figs.7-12. Calileuctra ephemera larval characters. 7. Lacinia, ventral. 8. Lacinia and galea, lateral. 9. Lacinia apex,

ventral. 10. Right mandible, lateral. 11. Left mandible, outside. 12. Left mandible outside molar cup teeth.

Arrows = molar cup.




Figs.13-18. Calileuctra ephemera larval characters. 13. Left mandible molar cup. 14. Left foreleg femur and tibia,

anterior. 15. Left foreleg tarsus, ventral. 16. Abdominal terga 8-10. 17. Abdominal tergum 7. 18. Abdominal

tergum 9.




Figs.19-20. Calileuctra ephemera cerci. 19. Basal 4 segments, dorsal. 20. Cercal segments 5, 6.

Calileuctra ephemera Shepard & Baumann

(Figs. 1-6, 7-12, 13-18, 19-20, 33)

Distribution and collection site descriptions.

Known from Putah Creek in Lake Co., tributaries

and the main channel of Whitlow Creek, Mendocino

Co., and the type locality, Capell Creek tributary in

Napa Co., all in California. Capell Creek tributary

and Putah Creek sites are both in the Putah Creek

drainage but separated by about 34 miles. The type

locality (Fig. 33) was described by Shepard &

Baumann (1995), whereas the latter site consists of

two small, adjacent stream channels; the larger,

southern channel, from which the female specimen

was collected, has a closed riparian canopy

composed primarily of California Bay (Umbellularia

californica (Hook & Arn.) Nutt.), Big-leaf Maple (Acer

macrophyllum Pursh), Blue Oak (Quercus douglasii

Hook & Arn.), Black Oak (Q. kelloggii Newberry),

Canyon Live Oak (Q. chrysolepis Liebm.) and

California Buckeye (Aesculus californica (Spach)

Nutt.). Additional shading occurs from the

surrounding mixed oak forest along adjacent steep

slopes. Herbaceous streamside vegetation consists

mostly of California Fescue (Festuca californica Vasey).

Adult Malenka depressa (Banks), Sweltsa pisteri

Baumann & Bottorff and Soliperla thyra (Needham &

Smith) were swept from riparian vegetation by

author Serpa, and larvae of Oemopteryx vanduzeea

(Claassen) and an undetermined Isoperla were

collected on other sample dates.

The Whitlow Creek tributary sites are located

about 43 miles northwest of the Putah Creek site (or

74 miles northwest of the type locality) on property

owned by the Conservation Fund in the Garcia River

watershed, Mendocino County, California, and the

site receives additional protection from an easement

held by the Nature Conservancy. The intermittent

tributaries flow through steep forested terrain,

becoming relatively level near their junctions with

the main channel of Whitlow Creek which retains

water at least 9 months of the year. The tributaries

flow for about 4-5 months of the year but probably

have hyporheic springs (suggested by the presence of

blind unpigmented amphipods, Stygobromus sp. and

the blind isopod, Callasellus californicus (Miller))

which reach the surface only during the wet months.

The portion of the forest in which Calileuctra

specimens were collected includes a heavy oak

component along with California Bay, Oregon Ash

(Fraxinus latifolia Benth.), Tan Oak (Notholithocarpus

densiflorus (Hook & Arn.) Manos, Cannon & S.H.Oh),

California Buckeye, Big-leaf Maple and young

Douglas fir. Oregon White Oak (Quercus garryana

Douglas ex. Hook.) grow on the adjacent slopes,

often extending over the stream and providing

additional shading. Seventy six adult C. ephemera

adults were swept or beat from riparian vegetation

with the vast majority of specimens found on

herbaceous Festuca, rushes or ferns. All larvae were




collected from pools where the sand-gravel substrate

could be disturbed to a depth of at least 3 inches.

Other stoneflies collected with Calileuctra include

members of the Capnia ventura Nelson & Baumann

complex (R.W. Baumann in litt.) and Mesocapnia

projecta (Frison).

Material examined. CALIFORNIA: Lake Co., Putah

Creek, Hwy 175, 3.5 mi N Middletown, 23.00 mile

marker, 17 May 1998, C.R. Nelson, B. Stark, S.W.

Szczytko, I. Sivec, 1♀ (C.R. Nelson Collection).

Mendocino Co., 15 collections by L.E. Serpa, 17-II-

2010 to 16-IV-2012, mainly tributary complex of

Whitlow Creek, 38°54’41.4” - 38°55’0’’N, 123°28’14” -

123°28’38”W, 131.4 - 204.5 m elevation, 33♂, 47♀, 15

larvae. One pair of larvae and most adults deposited

in L.E. Serpa collection, 1♂ and 2♀ deposited in R.W.

Baumann collection, Monte L. Bean Museum,

Brigham Young University, Provo, UT, and

remaining larvae and a few adults deposited in B.P.

Stark collection, Mississippi College, Clinton, MS.

Characters of mature larvae. Color, pigmentation

(Fig. 1), lacinia (Figs. 7, 8), general body, leg, and

cercal setation (Fig. 1,) and mouthparts, typical of

genus. Body length ♂ 5-6.5 mm, ♀ 6-8 mm. Head

capsule width ♂ 0.66-0.75 mm, ♀ 0.75-0.84 mm.

Antennal segments 60-64, each with very short apical

circlet of hairs or sensillae. Lacinia triangular and

palmate, with 2 rounded-tipped apical teeth, 8-10

dorsal and ventral rows of long, sharp spines, a long,

trichoid sensilum arising from the base of one apical

tooth (and as long as the tooth) (Fig. 9), and a

scalloped palm surface. Right mandibular molar cup,

in side view, with 8-10 stalked teeth that are

manifested in both Calileuctra species as stalked

ridges extending onto the cup (see inside surface of

C. dobryi cup, Fig. 27). Left mandibular molar cup

with about 12 long, curved, comb-like outer sharp

teeth, and 3 or 4 raised transverse ridges (that in Fig.

13 are probably well worn). Mesosternum with a

double stem of its Y-ridge (Fig. 2). Inside fore

wingpad length ♂ 0.66-0.75 mm, ♀ 0.72-0.96 mm;

inside hind wingpad length ♂ 0.51-0.63 mm, ♀ 0.60-

0.78 mm. Foreleg femoral and tibial surface with

sparse, short, stiff hairs, fine appressed clothing

hairs, and few if any fringe hairs (Figs. 1, 14); apex of

tibia with 2 heavy apical spines; tarsal segments 1, 2,

with double ventral rows of short spines (Fig. 15).

Abdominal terga with scattered, appressed clothing

hairs, and segments 7-10 with thick, bristle-like setae,

especially laterally, in dorsal view (Figs. 16-18); terga

7-8 (Fig. 17) without 2 diverging rows of short, thick

sensillae (as present in C. dobryi, see Figs. 30, 31).

Developing male genitalia evident dorsally and

ventrally just prior to emergence (Figs. 3, 4). Cercal

segments 20-24 (Fig. 6), each, except terminal few,

with apical circlet of more than 15 stiff hairs (Fig. 20);

hairs of basal segments only about half the length of

following segment (Fig. 19).

Calileuctra dobryi Shepard & Baumann

(Figs. 21-24, 25-30, 31-32, 34)

Distribution and collection site descriptions.

Known from two or possibly three sites in Los

Angeles County and two sites in Orange County in

Southern California. The Los Angeles County sites

include the type locality in South Fork Elsmere

Canyon, San Gabriel Mountains and East Fork

Arroyo Sequit, Santa Monica Mountains (Fig. 34). In

Orange County, Trabuco Canyon and Silverado

Canyon in the Santa Ana Mountains are the known

sites.

Material examined. Orange Co., 6 collections by

K.W. Stewart and E.F. Drake 6-IV-2004 to 22-II-2006,

same locality vicinity, Silverado Creek, Silverado

Canyon, at gravel low water crossing 161m above

Forest Boundary Gate, 33°44’55”N 117°34’58”W, 117♂,

19♀, 3 pairs in-copula, 4 larvae, 1 exuvium.

Deposited in B.P. Stark collection, Mississippi

College, Clinton MS.

Characters of mature larvae. (some measurements

and counts, as done for C. ephemera, were not made

because of the few larvae available for study). Color,

pigmentation, general body and cercal setation (Fig.

21), and mouthparts typical of genus. Body length 7-

8 mm. Head capsule width (1♀ larva) 0.75 mm.

Antennal segments 60-64, each with very short apical

circlet of hairs or sensillae. Lacinia triangulate and

palmate, with 2 rounded-tipped apical teeth, 8-10

dorsal and ventral rows of long, sharp spines (Fig.

25), and a scalloped palm surface. Right mandibular

molar cup outer margin, in side view, appears as a

row of blunt-tipped, stalked teeth (Fig. 26) that are

actually the outer edges of 10-12 stalked ridges

extending well onto the inner cup surface (Fig. 27).

Left mandible molar cup with comb-like outer teeth




Figs. 21-24. Calileuctra dobryi larval characters. 21. Habitus (from Stewart & Drake, 2007); scale line = 2mm.

22. Right foreleg. 23. Abdomen, left lateral (from Stewart & Drake, 2007). 24. Basal 16 cercal segments, dorsal

(from Stewart & Drake, 2007).

(Fig. 28; cup of the 1 larva sacrificed for SEM was

well worn, in poor condition). Mesosternum with a

double stem of its Y-ridge. Foreleg femoral and tibial

surface with sparse hairs and with variable number

of fine fringe hairs; apex of tibia with 2 heavy apical

spines (Fig. 22). Tarsal segments 1, 2 with double

vental row of short thick setae. Abdominal terga with

long, fine appressed surface clothing hairs; terga 7-9

without thick setae (Fig. 29), especially laterally, as

are present in C. ephemera (Figs. 16-18). Terga 7-8

21

22

23

24




Figs. 25-30. Calileuctra dobryi larval characters. 25. Right lacinia, ventral. 26. Right mandibular molar cup, outer

margin. 27. Ridges of right mandibular molar cup. 28. Worn left mandibular molar cup. 29. Abdominal

tergum 7, left side, arrow = sensilla row. 30. Abdominal tergum 8, diverging rows of short, thick sensillae.




Figs.31-32. Calileuctra dobryi larval characters. 31. Abdominal tergum 8, right side. 32. Basal cercal segments,

dorsal.

with 2 diverging basal, sublateral rows of short, thick

sensillae (Figs. 30-31), not present in C. ephemera.

Cercal segments 20-22, each with apical circlet of more

than 15 stiff hairs, except few apical and preapical

cercomeres (Fig. 16); basal segments with some hairs

longer than the following segment (Fig. 32).

Key to Mature Calileuctra larvae

1 Abdominal terga 7-10 with thick setae, especially

laterally Figs. 16-18); terga 7, 8 without diverging

sublateral rows of short, thick sensillae (Fig. 17).

Apical hair circlet on basal 3 cercal segments

mostly about 0.5-0.75 the length of following

segment (Fig. 19) ……...…………………. ephemera

1’ Abdominal terga 7-10 without thick setae (Fig.

29); terga 7, 8 with diverging sublateral rows of

short, thick sensillae (Figs. 30, 31). Some apical

circlet hairs on basal 3 cercal segments longer

than following segment (Fig. 33) ................... dobryi

GENERAL DISCUSSION

The now clarified structure of the molar cup of

the right mandible, with its stalked ridges forming a

masticating pad in C. dobryi (Fig. 27), and mentioned

in the written description of C. ephemera larvae by

Shepard & Baumann (1995), probably acts like a

“mortar” against the opposing, ridged molar cup

face (smooth and worn?, Fig. 13) of the left mandible

as a “pestle”. The long, curved comb-like teeth of the

outer edge of the left cup probably act to hold food

particles in the cup during mastication. Shepard &

Baumann (1995) indicated these molar areas of the

mandibles are similar to those of beetle larvae that

feed on fungal tissues with high protein content,

particularly of spores, that would aid in the fast

growth and development of larvae.

The original side-only views of the right larval

mandibles of leuctrid and capniid genera, drawn

from SEM’s by Stewart & Stark (2002), and described

by them as “scraping ridges” are now clarified in

Calileuctra, and in two species of Capnia (Stewart et al.

2011) more as masticating structures. The stalked

ridges of the right mandible of Calileuctra, and the

different, raised, bristly pads of Capnia umpqua

Frison 1942 and C. ventura Nelson & Baumann 1987

(see Figs. 46-47, 52-53 in Stewart et al. 2011) suggest

that further SEM study of larvae of additional genera

and species of these families may provide an

additional diagnostic character in the difficult

separation of their larvae. Current keys to larvae of

these two families use: 1) the difficult to observe

extent of the pleural fold on abdominal segments and

extent of coverage of the bases of the maxilla by the

mentum, and 2) inconsistent shapes of the abdominal

terga, length relationships of the wingpads, and




Figs. 33-34. Intermittent headwater habitats of Calileuctra. 33. Capell Creek, type locality of C. ephemera. 34.

Arroyo Sequit, habitat of C. dobryi. No larvae found in rainwater pools, February 2000.

inconsistent (for different leuctrid genera) presence

or absence of a posterior setal fringe on abdominal

terga (Stewart & Stark 2002, 2008).

The collections of adult Calileuctra of both species

indicate that, upon emergence, they remain in low

lying herbaceous vegetation, or objects close to the

stream. Good numbers of larvae have only been

collected by disturbing sand or sand-cobble

substrates within shallow pools, or slow current

reaches, of headwater tributaries in hills at elevations

near 130-140 meters.

ACKNOWLEDGEMENTS

This study was supported in part during earlier

years by grants from the University of North Texas

Faculty Research Fund. Bill Shepard, Cheryl Barr,

and Keith Dobry helped in collection attempts in

2000, and Eugene Drake helped with, and made

numerous collections of C. dobryi in Silverado

Canyon Creek, California, 2004-2006. We also thank

the Conservation Fund for their role in protecting the

Garcia River Forest, and for their gracious

cooperation with this and other scientific studies

conducted on their property. We also acknowledge

the long term efforts of the North Coast Regional

Water Quality Control Board to improve water

quality throughout the Garcia River watershed.

REFERENCES

Shepard, W.D. & R.W. Baumann. 1995. Calileuctra, a

new genus, and two new species of stoneflies

from California (Plecoptera: Leuctridae). Great

Basin Naturalist, 55:124-134.

Snellen, R.K. & K.W. Stewart. 1979. The life cycle and




drumming behavior of Zealeuctra claasseni

(Frison), and Zealeuctra hitei Ricker & Ross

(Plecoptera: Leuctridae) in Texas, USA. Aquatic

Insects, 1:65-89.

Stewart, K.W. & B.P. Stark. 2002. Nymphs of North

American stonefly genera (Plecoptera), 2nd ed. The

Caddis Press, Columbus, Ohio. 510 pp.

Stewart, K.W. & B.P. Stark. 2008. Plecoptera, Chapter

14. Pp. 311-384 in Merritt, R.W., K.W. Cummins &

M.B. Berg [editors]. An introduction to the

aquatic insects of North America, 4th ed.

Kendall/Hunt Publishing Company, Dubuque,

Iowa. 1158 pp.

Stewart, K.W. & E.F. Drake. 2007. The nymphs of two

rare western stonefly (Plecoptera) species,

representing little-known genera. Transactions of

the American Entomological Society, 133:115-122.

Stewart, K.W., E.F. Drake, & B.P. Stark. 2011. Larvae

of five species of the winter stonefly genus Capnia

(Plecoptera: Capniidae) from California, U.S.A.

Illiesia, 7:167-181.

Received 22 December 2012, Accepted 31 December 2012,

Published 9 January 2013

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